In 1926, the Florida State Road Department engaged
in making a road-cut immediately east of the Chipola River,
near Marianna, in Jackson County. During the course of
this operation a number of fossilized fish remains were
recovered, of which one was described as a new species of
Lutianus by Dr. William K. Gregory (1930). The unde-
scribed balance of the material was placed in storage,
where it remained until 1957, when the Florida Geological
Survey moved to new and permanent quarters. Amongthese
unidentified specimens has proved to be an example of a
beryciform fish.

The numerous fossil and living genera which are as-
signed to the Order Beryciformes (Berg, 1940), although of
widely variant structure, diverse adaptation, and consequently
complex taxonomy, are quite generally treated as a natural
assemblage. Great phylogenetic importance has been at-
tached to the group (Gregory, 1933; Regan, 1911; and Starks,
1904; among others) because of the common possession by
its members of a combination of characteristics immediate
between primitive "isospondyls and advanced spiny-rayed
teleosts. The beryciform fishes were well established and,
it is evident, of wide marine distribution around the world
by upper Cretaceous times. Notwithstanding, the details of

1Publishedwith the permission of the Secretary, Smith-
sonian Institution. 65369

1

FLORIDA GEOLOGICAL SURVEY

their origin and radiation remain obscure and, in fact, their
fossil record in the Western Hemisphere is negligible. It is
on this basis that the unique specimen from Florida warrants
the following description.

The authors acknowledge their appreciation of Dr.
Robert O. Vernon, Director, Florida Geological Survey,
for affording the opportunity of this study. Dr. Leonard P.
Schultz and staff, Division of Fishes, U. S. National Museum,
and Dr. P. E. Cloud, Jr., Dr. Roland W. Brown, and Miss
Ruth Todd, U. S. Geological Survey, have been most generous
of assistance and advice. Much of the compilation of the
comparative data employed has been carried out by Miss
Dianne Hubbard; and the illustrations have been prepared by
Mr. L. B. Isham and Mr. Jack Scott.

DESCRIPTION

The delicate specimen, identified by the Florida Geo-
logical Survey No. V-5776, and deriving from the lower part
of the Oligocene Marianna limestone, lacks parts anteriorly
and anteroventrally of the head, and of others dorsally along
the left flank. It, however, is preserved without either ap-
preciable compression or distortion and comprises about
two-thirds of an individual, which, when compared withholo-
centrid beryciformes, is of relatively elongate body habit.
The dorsum and venter are not longitudinally keeled, and in
transverse section the bodyis of regularly ovate outline. On
the basis of comparison with the living Myripristis, the fol-
lowing dimensions in millimeters for the fossil have been
estimated: standard length, 250; head length from rostral
tip to posterior border of opercular spine, 74; and maximum
body depth (not exceeding), 96.

The skull is indicated to have been somewhat longer
than deep. If the above approximations of size are correct,
the head, relative to the standard length, is small; and in this
character the fossil compares more closely to Holocentrus
and Myripristis than to other related Recent genera. Impres-
sion on the matrix of the orbital surfaces of the parethmoids
denotes abreadth of snout fully equal to the postorbital width

2

SPECIAL PUBLICATION NO. 2 PAPER NO. 3

of the skull (fig. 1).

Large frontals cover practically the entire, trans-
versely convex skull roof. The elements are joined in a
median longitudinal suture whichis depressedinto a groove.
Strong ridges bearing a fewposteriorly radiating and dentate
ribs and moderately deep mucous channels mark the dorsal
surfaces of the bones. The parietals are small, posteriorly
restricted, and widely separated from each other. Only
narrow margins of the sphenotics, pterotics, epiotics and
supra-occipital could have been exposed laterally and pos-
teriorly in any dorsal view of the skull.

The occipital face of the neurocranium extends down-
ward and backward from the posterior edge of the skull roof
in an angle of approximately 45 from the horizontal (fig. 2).
The skullis thus prolonged in a spino-occipital development
greater than that displayed by the skulls of living relatives
with which comparison has been made. The supra-occipital
crest is produced backwardlyandis neither elevated dorsally
nor extended anteriorly in the midline of the roof. This fact,
coupled with evidences of deep and completely roofed post-
temporal fossae, suggests strongly that the cranial insertions
of the epaxial body musculature were confinedto the posterior
surface of the skull. Insofar as canbe observed, the foramen
magnum is bounded ventrally by a vertebra which is firmly
ankylosed to the more anteriorly lying basi-occipital. The
exoccipitals, which do not contribute to the formation of the
occipital condyle, form the lateral margins of the same
endocranial opening and if, in fact, they do not meet each
other dorsally in the midline, only the narrow inferior ex-
tremity of the supra-occipital spine can intervene. The
ventrolateral extent of the exoccipitals is unknown. How-
ever, in dorsal aspect, because of the spino-occipital devel-
opment of the skull, the posteroventral parts of each lie
horizontally on the underlying bones and appear narrow and
attenuated. Anteriorly the exoccipitals gradually widen and
flare dorsally in a concave posterolaterally directed face.
Remains of both epiotic elements are preserved. Those on
the right side comprise the broad, triangularly expanded,
post-temporal process and the more anterior dorsolateral
lamina, which meeting the dorsal wing of the pterotic, form
the posterior portion of the roof of the post-temporalfossa.
The fragment of the left epiotic is that anterovertical part
constituting the mesial wall of the left post-temporal fossa.

The condition of this remnant and inclination of adjacent
exoccipital part indicate that the fossae opened backwardly
with sharp mesial and ventral lips. In all the living holo-
centrids examined, the fossae are posteriorly deepened and
their floors are continuedas grooves, convergent toward the
foramen magnum, across the posterodorsal faces of the
exoccipitals (fig. 3).

The exposure of the intercalar (opisthotic of Nelson,
1957)onthe posterior face of the skull is limited tothe proc-
ess serving the ventral point of attachment for the post-
temporal bone. This is apparently situated belowthe lateral
third of the width of the post-temporal fossa. The element
does not possess a mesially directed splint applied to the
exoccipital. Laterallythe intercalar extends forwardacross
fully one-half, the length of the otic region, flooring a deep
concavity immediately below the articular facet of the
hyomandibular. Similar but less well developed depressions
in Holocentrus and Myripristis display in their anterior parts
the posterior opening of the jugular canal and the foramen for
the glossopharyngeal nerve, and their anterior and dorsal
walls serve as areas of origin for at least the adductor
hyomandibularis and adductor operculi muscles.

The dorsally and anteriorly succeeding pterotic,
sphenotic, pro-otic, and alisphenoid bones in the specimen
are either damaged or poorly exposed. However, the dorso-
lateral flange of the pterotic and the high anterodor sal lamina
of the sphenotic encompass a relatively large, lateral tem-
poral fossa which provides origin for the dilator operculi
and elevator palatine muscles. As the region is exposed on the
left side, the lateral wall of the jugular canal and trigemino-
facialis chamber is missing, but the pro-otic is inflated
posteroventrally and forms a portion of the anterior wall of
the auditory bulla. The out-turned lip of this part indicates
a flat, laterally facing, membraneous area external to the
saccular region.

Two elements of the infra-orbital series of bones are
preserved intact on the right side (fig. 4). These, situated
below the orbit, are relativelylarge and dividedinto shallow
external and deep internal lamina by the anteroposterior
passage of the broad infra-orbital mucous channel across

SPECIAL PUBLICATION NO. Z PAPER NO. 3

their surfaces. The margins of these laminae are only
weakly dentate.

Of the visceral skeleton, the possession of two supra-
maxillary bones is shown. Although lying in approximately
correct position on the left side of the specimen, it must be
presumed that these elements are displaced from tne right
and as thus interpreted display the same configurations and
relationships as the corresponding supra-maxillaries of
Myripristis. The hyomandibula has double articular heads
and the thin anterior lamella of the bone is thus fairlybroad.
The characteristic external rib of the bone is well developed,
and situated near the posterior border of the bone. The
vertical portion of this rib below the level of the opercular
process projects laterally in a practically true transverse
plane. Canals for the passage of branches of the hyomandi-
bular nerve emerge both dorsally and anteriorly to this ridge.

The preoperculum is a short, exceedingly deep crescent
of bone which displays neither a sharp forward angulation of
its interior part nor a posterior spine. The thickness of the
anterior border of the bone compliments the transverse in-
clination of the vertical hyomandibular rib andthe posterior
margins of the laminae bounding the vertical preopercular
mucous channel are finely dentate throughout their extent.
A single vertical series of scales underlies the posterior
margin of the preoperculum and overlies the anterior portion
of the operculum. The longitudinally ornamented expanse
of the operculum behind these scales is deeper than long
with generally concave, dentate margins above, and below a
posterior spinous process.

The remains ofat least eight spinous rays of the dorsal
fin are to be observed. Of these the foremost one preserved
is the largest. These facts denote a long-based spinous
dorsal fin composed of 10 or more rays originating above or
slightly behind the posterior margin of the operculum. As
in the Recent Myripristis, the individual spines are bilater-
ally asymmetrical and in a retractedposition, and alternate
with each other in lying first on one side of the median line
and then the other.

Scalation was apparently complete over the body and
cheek. The scales, although somewhat variable in size

regionally, are generally large. Eight occur in the vertical
rows between the venter and longitudinal lateral line sequence
below the spinous dorsal fin. Individual examples from the
anterior flank measure approximately 17. 5 millimeters deep
by 12. 0 millimeters long. In outline, dorsal, posterior, and
ventral borders are confluentandperform a broadly rounded
curve. The basal margin is of low forward convexity, and
in consequence the dorsal and ventral basal angles are well
defined. The nucleus is a small, vertically ovate locus sit-
uated slightly nearer the dorsal and apical margins of the
scales than to the ventral and basal ones. On anterior body
scales the nuclear fields are unornamented. Posteriorly
they appear larger in proportion to total scale area and may
be markedby moderately coar se and entirely vertical circuli.
Surrounding the nuclei, fine, crescentically arranged, and
weakly incised circuli cover the dorsal, basal, and ventral
quadrants of the scale. All appear to parallel the scale mar-
gins. No basal radii are present but suggestive of these are
five or six regular series ofplications in the courses of the
circuli, which diverge fromthe nuclear area across the em-
bedded anterior scale part. Posteriorly most of the circuli
stop along the apical diagonal. However, one of every third
or fourth crosses these diagonals onto the apical field where
they curve obliquely away from the longitudinal axis of the
scale toward the apical periphery in radial fashion. The
posteriorly exposed apical field displays a nuclear angle of
nearly 180. Parallel and longitudinally directed grooves
(numbering about four in one millimeter) traverse the ex-
ternal surface of the part. The ribs between the grooves
project as teeth along the posterior border of the scale and
their sides may be obliquely striated.

REMARKS

The presently described fish must be considered, at
this time, of the same identity as the single holocentridfrom
the Ocala limestone of Jackson County, Florida, described
by Conrad (1941) under the name Holocentrites ovalis. To
be sure, certain apparent variations are to be observed be-
tween the Oligocene specimen and the Eocene genoholotype.
However, no attempt to determine the validity and significance
of these seems feasible. It is only with the greatest diffi -
culty that the exact kind and degree of variation between

14

SPECIAL PUBLICATION NO. 2 PAPER NO. 3 15

incompletely and differently preserved specimens can be
established. In any event, onthebasis of current knowledge,
Holocentrites is distinguishable from all other fossil and
Recent genera of the family Holocentridae (Arambourg, 1927;
Bertin and Arambourg, 1958; Nelson, 1955; Pauca, 1931;
Romer, 1945; Starks, 1904; Woods, 1955; Woodward, 1901
and 1902; etc. ). Further, the new information afforded by
the Oligocene specimen permits revision of the diagnosis of
this genus, as follows: A relatively fusiform holocentrid
with a maximum body length of about one-fourth the total
length a vertebral formula of 11 + 14 + 1 = 26. The fish
resembles Myripristis in the strength of the principal and
fewness of the radiating frontal ridges, the correspondingly
good definition of the mucous channels, the expanded post-
temporal process of the epiotic, the protuberant lips of the
laterally directed membranous face of the auditory bulla,
the deeply channeled and weakly ornamented infra-orbital
bones, and the simple crescentic and nonspinous preoper-
culum. It differs from Myripristis and is similar to Holo-
centrus in possessing abroad, double-headed hyomandibula;
but is distinct from the two noted living genera by virtue of
a relatively greater spino-occipital development and fusion
of a vertebra to the basi-occipital, which prevents the two
exoccipitals meeting each other in the midline below the
foramen magnum.

The Holocentridae date from the early upper Creta-
ceous when such referred genera as Homonotichthys,
Trachichthyoides, and Caproberyx, from Turonian horizons
in England, first appear in the fossil record. Occurrence
of family representatives during succeeding geologic periods
to the present time are not frequent but indicate a continuous
existence. A more complete representation of the group,
the detailed study of these and available specimens, and
recognition of either morphologically or temporally older
antecedents would facilitate reconstruction of the phylogenetic
history of the family. In the absence of such objective data,
however, recourse must be taken in the analysis of certain
obvious osteological characteristics selected because of
availability to examination on paleontological materials.
Two principal adaptive conditions are apparent among living
holocentrids. The first of these, exemplifiedby Myripristis,
combines well developed mucous channels; longitudinal

FLORIDA GEOLOGICAL SURVEY

crests on the frontals of moderate width and bearing few
radiating ribs; a nonspinous, crescentic preoperculum;
short, high, and single headed hyomandibula; and "normal"
premaxillary. The second condition, characterized by
Holocentrus, displays much reduced mucous channels;
principal frontal crests very broad with numerous radiating
ribs; a spinous, sharply angulated preoperculum; broad,
double-headed hyomandibula; and premaxillary with highly
developed ascending process. While, as implied in the
literature, Holocentrus may be more advanced toward the
"true percoids" than Myripristis, by reason of the spinous
preoperculum and more protrusile premaxillary, the two
adaptations can be traced without question back through the
Tertiary into the Eocene. The upper Cretaceous holocentrid
genera all possess the simply crescentic andnonspinous pre-
operculum; broad, double-headed hyomandibula; and pre-
maxillary with low ascending processes. The phylogeny of
the Holocentridae consequently has been pictured (Conrad,
1941) as an early Tertiary divergence of the myripristine
and holocentrine lineages from a single holocentrid stem of
Cretaceous origin.

Homonotichthys is too poorly known for comment, but
in degree of development of mucous channels and frontal
crests Trachichthyoides and Holocentrites are not radically
different from Myripristis, and Caproberyx would seem to
approach Holocentrus. Development of the frontal bones may
be far too inadequate a criterionbut, in contrast to the above
phylogenetic concept, a basis is suggested for recognizing
the distinctiveness of the two "subfamilial" groups for the
entire known history of the family. By this latter view, the
single-headed hyomandibulae of Myripristis and the more
protrusile premaxilli of Holocentrus would be regarded as
adaptive morphologic distinctions of the visceral skeleton
whose selective values were established only during post-
Eocene times. Some degree of support would seem given to
this opinion (oral communication from W. A. Gosline) by the
respective habits of Myripristis and Holocentrus. The un-
doubted flexibility of the entire visceral complex resulting
from a single-headed hyomandibula would be mechanically
advantageous to Myripristis, which feeds in middle and upper
depths of water by "sifting "pelagic zooplankton through long
gill rakers. Conversely, Holocentrus utilizes protrusile

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SPECIAL PUBLICATION NO. 2 PAPER NO. 3 17

premaxilli for 'browsing" on benthonic animals.

Fossil fishes in general are of rare occurrence in
nature. Corroborative experimental data have not been found
but it has been widely assumed thatthe rate of decomposition
of the dead bodies of these animals is comparatively rapid
under aerobic conditions. Conversely, while not indicative
of life habitats, the best preserved skeletons are generally
attributed to reduction environments (cf. Rayner, 1958).

The Marianna limestone, of lower Oligocene age, has
been described (Moore, 1955) as a 15-foot thick layer of
soft, cream to white, massive limestone. It has a limited
distribution, outcropping over most of Jackson County,
Florida, and occurring inthe subsurface only as far west as
Walton County. Puri and Vernon (1956) refer to the presence
in the formation of abundant Lepidocyclina, as index fossils.
From the matrix removed from the presently described
specimen during preparation, Miss Ruth Todd, of the U. S.
Geological Survey, has picked and identified the following
list of microfossils:

About these Miss Todd has stated (in littoris): "Four
features of this foraminiferal assemblage point to its having
been deposited in moderately deep water (such as 10 fathoms
or more): (a)presence of globigerinids; (b)type of benthonic
genera, Recent representatives of which are more likely to
occur in deep than in shallow water (Robulus, Nodosaria,
Uvigerina, Angulogerina, Eponides); (c) absence of miliolids

FLORIDA GEOLOGICAL SURVEY

usually considered to indicate shallow deposition; and (d) thin-
walled nature of the tests in contrast to the heavy-walled tests
commonly found in shallow water under the effect of wave
motion. "

To be pointed out also is the fact that the foraminif-
eral assemblage undoubtedly indicates well oxygenatedbottom
water. Under these circumstances the geological environ-
ment of the originally well preserved specimens of Holo-
centrites and associated fishes is of considerable interest.
One plausible explanation was suggested by Dr. Preston E.
Cloud, Jr. (oral communication), who called the attention
of these writers to the relationship of sediment particle size
to density of anaerobic bacterial populations below the sedi-
ment water interface and the amount of gaseous interchange
between water and the substratum. Entombment of these
fishes in a sediment as fine-grained as the Marianna lime-
stone might well insure anaerobic preservation. Recent
squirrel fishes are said (Longley and Hildebrand, 1941) to
secrete themselves in practically any available crevice or
depression by day and to forage for food by night. The snap-
pers also are reported(ibidem) of somewhat similar habits,
idling in schools in protected situations during the day and
feeding singly at night. If similar behavioral patterns were
displayed by the fossil representatives of these two groups,
the chances of burial, although perhaps accidental in every
case, would be much enhanced.